Respiration rates and abundances of dominant copepods of the northern Benguela Current System

Respiration rates of 16 calanoid copepod species from the northern Benguela upwelling system were measured on board RRS Discovery in September/October 2010 to determine their energy requirements and assess their significance in the carbon cycle. Individual respiration rates were standardised to a mean copepod body mass and a temperature regime typical of the northern Benguela Current. These adjusted respiration rates revealed two different activity levels (active and resting) in copepodids C5 of Calanoides carinatus and females of Rhincalanus nasutus, which reduced their metabolism during dormancy by 82% and 62%, respectively. An allometric function (Imax) and an energy budget approach were performed to calculate ingestion rates. Imax generally overestimated the ingestion rates derived from the energy budget approach by >75%. We suggest that the energy budget approach is the more reliable approximation with a total calanoid copepod (mainly females) consumption of 78 mg C m-2 d-1 in neritic regions and 21 mg C m-2 d-1 in oceanic regions. The two primarily herbivorous copepods C. carinatus (neritic) and Nannocalanus minor (oceanic) contributed 83% and 5%, respectively, to total consumption by calanoid copepods. Locally, C. carinatus can remove up to 90% of the diatom biomass daily. In contrast, the maximum daily removal of dinoflagellate biomass by N. minor was 9%. These estimates imply that C. carinatus is an important primary consumers in the neritic province of the northern Benguela system, while N. minor has little grazing impact on phytoplankton populations further offshore. Data on energy requirements and total consumption rates of dominant calanoid copepods of this study are essential for the development of realistic carbon budgets and food-web models for the northern Benguela upwelling system.

Solar radiation over and under sea ice during POLARSTERN cruise PS92 (TRANSSIZ) in 2015

Measurements of solar radiation over and under sea ice have been performed on various stations in the Arctic Ocean during the Polarstern cruise PS92 (TRANSSIZ) between 19 May and 30 June 2015. All radiation measurements have been performed with Ramses spectral radiometers (Trios, Rastede, Germany). All data are given in full spectral resolution interpolated to 1.0 nm, and integrated over the entire wavelength range (broadband, total: 320 to 950 nm). Two sensors were mounted on a Remotely Operated Vehicle (ROV) and one radiometer was installed on the sea ice for surface reference measurements (solar irradiance). On the ROV, one irradiance sensor (cos-collector) for energy budget calculations and one radiance sensor (9° opening angle) to obtain high resolution spatial variability were installed. Along with the radiation measurements, ROV positions were obtained from acoustic USBL-positioning and all parameters of vehicle depth, distance to the ice and attitude recorded. All times are given in UTC.

Bioenergetic trade-offs in the sea cucumber Apostichopus japonicus (Echinodermata: Holothuroidea) in response to CO2-driven ocean acidification

Ocean acidification (OA) caused by excessive CO2 is a potential ecological threat to marine organisms. The impacts of OA on echinoderms are well-documented, but there has been a strong bias towards sea urchins, and limited information is available on sea cucumbers. This work examined the effect of medium-term (60 days) exposure to three pH levels (pH 8.06, 7.72, and 7.41, covering present and future pH variability) on the bioenergetic responses of the sea cucumber, Apostichopus japonicus, an ecologically and economically important holothurian in Asian coasts. Results showed that the measured specific growth rate linearly decreased with decreased pH, leading to a 0.42 %/day decrease at pH 7.41 compared with that at pH 8.06. The impacts of pH on physiological energetics were variable: measured energy consumption and defecation rates linearly decreased with decreased pH, whereas maintenance energy in calculated respiration and excretion were not significantly affected. No shift in energy allocation pattern was observed in A. japonicus upon exposure to pH 7.72 compared with pH 8.06. However, a significant shift in energy budget occurred upon exposure to pH 7.41, leading to decreased energy intake and increased percentage of energy that was lost in feces, thereby resulting in a significantly lowered allocation into somatic growth. These findings indicate that adult A. japonicus is resilient to the OA scenario at the end of the twenty-first century, but further acidification may negatively influence the grazing capability and growth, thereby influencing its ecological functioning as an "ecosystem engineer" and potentially harming its culture output.

Seawater carbonate chemistry and resource allocation and extracellular acid-base status in the sea urchin Strongylocentrotus droebachiensis during experiments, 2012

Anthropogenic CO2 emission will lead to an increase in seawater pCO2 of up to 80-100 Pa (800-1000 µatm) within this century and to an acidification of the oceans. Green sea urchins (Strongylocentrotus droebachiensis) occurring in Kattegat experience seasonal hypercapnic and hypoxic conditions already today. Thus, anthropogenic CO2 emissions will add up to existing values and will lead to even higher pCO2 values >200 Pa (>2000 µatm). To estimate the green sea urchins' potential to acclimate to acidified seawater, we calculated an energy budget and determined the extracellular acid base status of adult S. droebachiensis exposed to moderately (102 to 145 Pa, 1007 to 1431 µatm) and highly (284 to 385 Pa, 2800 to 3800 µatm) elevated seawater pCO2 for 10 and 45 days. A 45 - day exposure to elevated pCO2 resulted in a shift in energy budgets, leading to reduced somatic and reproductive growth. Metabolic rates were not significantly affected, but ammonium excretion increased in response to elevated pCO2. This led to decreased O:N ratios. These findings suggest that protein metabolism is possibly enhanced under elevated pCO2 in order to support ion homeostasis by increasing net acid extrusion. The perivisceral coelomic fluid acid-base status revealed that S. droebachiensis is able to fully (intermediate pCO2) or partially (high pCO2) compensate extracellular pH (pHe) changes by accumulation of bicarbonate (maximum increases 2.5 mM), albeit at a slower rate than typically observed in other taxa (10 day duration for full pHe compensation). At intermediate pCO2, sea urchins were able to maintain fully compensated pHe for 45 days. Sea urchins from the higher pCO2 treatment could be divided into two groups following medium-term acclimation: one group of experimental animals (29%) contained remnants of food in their digestive system and maintained partially compensated pHe (+2.3 mM HCO3), while the other group (71%) exhibited an empty digestive system and a severe metabolic acidosis (-0.5 pH units, -2.4 mM HCO3). There was no difference in mortality between the three pCO2 treatments. The results of this study suggest that S. droebachiensis occurring in the Kattegat might be pre-adapted to hypercapnia due to natural variability in pCO2 in its habitat. We show for the first time that some echinoderm species can actively compensate extracellular pH. Seawater pCO2 values of >200 Pa, which will occur in the Kattegat within this century during seasonal hypoxic events, can possibly only be endured for a short time period of a few weeks. Increases in anthropogenic CO2 emissions and leakages from potential sub-seabed CO2 storage (CCS) sites thus impose a threat to the ecologically and economically important species S. droebachiensis.

Solar radiation over and under sea ice at ROV station PS86/081-1 during POLARSTERN cruise PS86 (AURORA) in July 2014

The observed changes in physical properties of sea ice such as decreased thickness and increased melt pond cover severely impact the energy budget of Arctic sea ice. Increased light transmission leads to increased deposition of solar energy in the upper ocean and thus plays a crucial role for amount and timing of sea-ice-melt and under-ice primary production. Recent developments in underwater technology provide new opportunities to study light transmission below the largely inaccessible underside of sea ice. We measured spectral under-ice radiance and irradiance using the new Nereid Under-Ice (NUI) underwater robotic vehicle, during a cruise of the R/V Polarstern to 83°N 6°W in the Arctic Ocean in July 2014. NUI is a next generation hybrid remotely operated vehicle (H-ROV) designed for both remotely piloted and autonomous surveys underneath land-fast and moving sea ice. Here we present results from one of the first comprehensive scientific dives of NUI employing its interdisciplinary sensor suite. We combine under-ice optical measurements with three dimensional under-ice topography (multibeam sonar) and aerial images of the surface conditions. We investigate the influence of spatially varying ice-thickness and surface properties on the spatial variability of light transmittance during summer. Our results show that surface properties such as melt ponds dominate the spatial distribution of the under-ice light field on small scales (<1000 m**2), while sea ice-thickness is the most important predictor for light transmission on larger scales. In addition, we propose the use of an algorithm to obtain histograms of light transmission from distributions of sea ice thickness and surface albedo.

Phenotypic plasticity of coralline algae in a High CO2 world

It is important to understand how marine calcifying organisms may acclimatize to ocean acidification to assess their survival over the coming century. We cultured the cold water coralline algae, Lithothamnion glaciale, under elevated pCO2 (408, 566, 770, and 1024 µatm) for 10 months. The results show that the cell (inter and intra) wall thickness is maintained, but there is a reduction in growth rate (linear extension) at all elevated pCO2. Furthermore a decrease in Mg content at the two highest CO2 treatments was observed. Comparison between our data and that at 3 months from the same long-term experiment shows that the acclimation differs over time since at 3 months, the samples cultured under high pCO2 showed a reduction in the cell (inter and intra) wall thickness but a maintained growth rate. This suggests a reallocation of the energy budget between 3 and 10 months and highlights the high degree plasticity that is present. This might provide a selective advantage in future high CO2 world.

Effects of seawater pCO2 and temperature on shell growth, shell stability, condition and cellular stress of Western Baltic Sea Mytilus edulis (L.) and Arctica islandica (L.)

Acidification of the World's oceans may directly impact reproduction, performance and shell formation of marine calcifying organisms. In addition, since shell production is costly and stress in general draws on an organism's energy budget, shell growth and stability of bivalves should indirectly be affected by environmental stress. The aim of this study was to investigate whether a combination of warming and acidification leads to increased physiological stress (lipofuscin accumulation and mortality) and affects the performance [shell growth, shell breaking force, condition index (Ci)] of young Mytilus edulis and Arctica islandica from the Baltic Sea. We cultured the bivalves in a fully-crossed 2-factorial experimental setup (seawater (sw) pCO2 levels "low", "medium" and "high" for both species, temperature levels 7.5, 10, 16, 20 and 25 °C for M. edulis and 7.5, 10 and 16 °C for A. islandica) for 13 weeks in summer. Mytilus edulis and A. islandica appeared to tolerate wide ranges of sw temperature and pCO2. Lipofuscin accumulation of M. edulis increased with temperature while the Ci decreased, but shell growth of the mussels only sharply decreased while its mortality increased between 20 and 25 °C. In A. islandica, lipofuscin accumulation increased with temperature, whereas the Ci, shell growth and shell breaking force decreased. The pCO2 treatment had only marginal effects on the measured parameters of both bivalve species. Shell growth of both bivalve species was not impaired by under-saturation of the sea water with respect to aragonite and calcite. Furthermore, independently of water temperatures shell breaking force of both species and shell growth of A. islandica remained unaffected by the applied elevated sw pCO2 for several months. Only at the highest temperature (25 °C), growth arrest of M. edulis was recorded at the high sw pCO2 treatment and the Ci of M. edulis was slightly higher at the medium sw pCO2 treatment than at the low and high sw pCO2 treatments. The only effect of elevated sw pCO2 on A. islandica was an increase in lipofuscin accumulation at the high sw pCO2 treatment compared to the medium sw pCO2 treatment. Our results show that, despite this robustness, growth of both M. edulis and A. islandica can be reduced if sw temperatures remain high for several weeks in summer. As large body size constitutes an escape from crab and sea star predation, this can make bivalves presumably more vulnerable to predation with possible negative consequences on population growth. In M. edulis, but not in A. islandica, this effect is amplified by elevated sw pCO2. We follow that combined effects of elevated sw pCO2 and ocean warming might cause shifts in future Western Baltic Sea community structures and ecosystem services; however, only if predators or other interacting species do not suffer as strong from these stressors.

Temperature and acidification variability reduce physiological performance in the intertidal zone porcelain crab Petrolisthes//cinctipes

We show here that increased variability of temperature and pH synergistically negatively affects the energetics of intertidal zone crabs. Under future climate scenarios, coastal ecosystems are projected to have increased extremes of low tide-associated thermal stress and ocean acidification-associated low pH, the individual or interactive effects of which have yet to be determined. To characterize energetic consequences of exposure to increased variability of pH and temperature, we exposed porcelain crabs, Petrolisthes cinctipes, to conditions that simulated current and future intertidal zone thermal and pH environments. During the daily low tide, specimens were exposed to no, moderate or extreme heating, and during the daily high tide experienced no, moderate or extreme acidification. Respiration rate and cardiac thermal limits were assessed following 2.5 weeks of acclimation. Thermal variation had a larger overall effect than pH variation, though there was an interactive effect between the two environmental drivers. Under the most extreme temperature and pH combination, respiration rate decreased while heat tolerance increased, indicating a smaller overall aerobic energy budget (i.e. a reduced O2 consumption rate) of which a larger portion is devoted to basal maintenance (i.e. greater thermal tolerance indicating induction of the cellular stress response). These results suggest the potential for negative long-term ecological consequences for intertidal ectotherms exposed to increased extremes in pH and temperature due to reduced energy for behavior and reproduction.

Solar radiation over and under sea ice during the POLARSTERN cruise ARK-XXVI/3 (TransArc) in summer 2011

Arctic sea ice has declined and become thinner and younger (more seasonal) during the last decade. One consequence of this is that the surface energy budget of the Arctic Ocean is changing. While the role of surface albedo has been studied intensively, it is still widely unknown how much light penetrates through sea ice into the upper ocean, affecting sea-ice mass balance, ecosystems, and geochemical processes. Here we present the first large-scale under-ice light measurements, operating spectral radiometers on a remotely operated vehicle (ROV) under Arctic sea ice in summer. This data set is used to produce an Arctic-wide map of light distribution under summer sea ice. Our results show that transmittance through first-year ice (FYI, 0.11) was almost three times larger than through multi-year ice (MYI, 0.04), and that this is mostly caused by the larger melt-pond coverage of FYI (42 vs. 23%). Also energy absorption was 50% larger in FYI than in MYI. Thus, a continuation of the observed sea-ice changes will increase the amount of light penetrating into the Arctic Ocean, enhancing sea-ice melt and affecting sea-ice and upper-ocean ecosystems.

The distribution of macrobenthos in surface sediments at Fladen Ground stations, North Sea

A general study of structure, biomass estimates and dynamics on the macrofauna was carried out in August 1975 and March 1976 during PREFLEX (1975) and FLEX (1976), the Fladen Ground Experiment. On the basis of these data an attempt was made to estimate macrobenthic production expressed as minimum production (MP). The macrobenthic production is discussed together with meiobenthic annual production and with indirectly estimated microbenthic production in relation to an energy input from the water column of about 25 g C m**-2 year**-1. From the production estimates of the three benthic components a rough energy budget is proposed. Sampling was performed at five stations for endofauna twice during the time of investigation and for epifauna once. At each station two replicate box core samples (30 X 20 cm) were taken for endofauna. Epifauna was sampled with an Agassiz trawl once at each station. The total numbers of endofauna increased from station 1 to 5. This was valid as well for August 1975 (4,233-12,166 individuals per m**2 and 10 cm sediment depth) as for March 1976 (1,008-2,925 individuals). The polychaetes were the dominant organisms with a share of 33 to 62 %. The densities for the endofauna decreased from August 1975 to March 1976 by a mean factor of 2.8. Abundances of epifauna amounted to values between 11 and 102 individuals per 1000 m**2. The biomass dry weights (DWT) for macrobenthic endofauna varied between 0.97 g DWT m**-2 and 6.42 g DWT m**-2 in August 1975 and between 0.27 g DWT m**-2 and 2.64 g DWT m**-2 in March 1976. The mean amounted to 1.74 g DWT m**-2. Dry weights of epifauna biomass gave values between 4.9 and 83.1 g DWT * 1000 m**-2. The minimum production for the total macro-endofauna at Fladen Ground amounted to 1.43 g DWT m**-2 yr**-1 or 0.82 g C m**-2 yr**-1. This resulted in a minimum turnover rate (P/B) of 0.8. The share produced by the polychaetes amounted to 1.06g DWT m**-2 yr**-1 or 74 %.

Rainforests at the beginning of the 21st century

Rainforests are situated at low latitude where forests enjoy steady and strong radiation. Biodiversity in rainforests has been very high, for historical and climatic reasons. The number of species is very high and tends to increase with precipitation and decrease with seasonality. Disturbance, soil fertility and forest stature also influence the species richness and high turnover of species contribute to diversity. Field observation and studies revealed that large scale deforestation could alter the regional and global climate significantly. Deforestation alters the surface albedo which leads to climate change. Regional land use contributes to climate change through surface-energy budget, as well as the carbon cycle. Forest fragmentation, logging, overhunting, fire and the expanding agriculture threaten the biodiversity. Rainforest covered area has significantly shrunk in the last decades. It is hard to protect the forests because of the growing demand for agricultural area and forest-derived products. Most measures proved ineffective to slow down the destruction. Hence, more forest will be lost in the future. Conservationists should take into consideration the secondary forests because biodiversity can be high enough and it is worth protecting them.